functionally-altered structures in domestic animals. The hypothesis of
panmixia is, as we have seen, out of court; and therefore Mr. Darwin's
groups of evidences are reinstated. There is the changed ratio of
wing-bones and leg-bones in the duck; there are the drooping ears of cats
in China, of horses in Russia, of sheep in Italy, of guinea-pigs in
Germany, of goats and cattle in India, of rabbits, pigs, and dogs in all
long-civilized countries. Though artificial selection has come into play
where drooping has become a curious trait (as in rabbits), and has probably
caused the greater size of ears which has in some cases gone along with
diminished muscular power over them; yet it could not have been the
initiator, and has not been operative on animals bred for profit. Again
there are the changes produced by climate; as instance, among plants, the
several varieties of maize established in Germany and transformed in the
course of a few generations.
Facts of another class are yielded by the blind inhabitants of caverns. One
who studies the memoir by Mr. Packard on _The Cave Fauna of North America_,
&c., will be astonished at the variety of types in which degeneration or
loss of the eyes has become a concomitant of life passed in darkness. A
great increase in the force of this evidence will be recognized on learning
that absence or extreme imperfection of visual organs is found also in
creatures living in perpetual night at the bottoms of deep oceans.
Endeavours to account for these facts otherwise than by the effects of
disuse we have seen to be futile.
Kindred evidence is yielded by decrease of the jaws in those races which
have had diminished use of them--mankind and certain domestic animals.
Relative smallness in the jaws of civilized men, manifest enough on
comparison, has been proved by direct measurement. In pet dogs--pugs,
household spaniels--we find associated the same cause with the same effect.
Though there has been artificial selection, yet this did not operate until
the diminution had become manifest. Moreover there has been diminution of
the other structures concerned in biting: there are smaller muscles, feeble
zygomata, and diminished areas for insertion of muscles--traits which
cannot have resulted from selection, since they are invisible in the living
animal.
In abnormal vision produced by abnormal use of the eyes we have evidence of
another kind. That the Germans, among whom congenital short sight is
notoriously prevalent, have been made shortsighted by inheritance of
modifications due to continual reading of print requiring close attention,
is by some disputed. It is strange, however, that if there exists no causal
connexion between them, neither trait occurs without the other elsewhere.
But for the belief that there is a causal connexion we have the verifying
testimony of oculists. From Dr. Lindsay Johnson I have cited cases within
his professional experience of functionally-produced myopia transmitted to
children; and he asserts that other oculists have had like experiences.
Development of the musical faculty in the successive members of families
from which the great composers have come, as well as in the civilized races
at large, is not to be explained by natural selection. Even when it is
great, the musical faculty has not a life-saving efficiency as compared
with the average of faculties; for the most highly gifted have commonly
passed less prosperous lives and left fewer offspring than have those
possessed of ordinary abilities. Still less can it be said that the musical
faculty in mankind at large has been developed by survival of the fittest.
No one will assert that men in general have been enabled to survive and
propagate in proportion as their musical appreciation was great.
The transmission of nervous peculiarities functionally produced is alleged
by the highest authorities--Dr. Savage, president of the Neurological
Society, and Dr. Hughlings Jackson. The evidence they assign confirms, and
is confirmed by, that which the development of the musical faculty above
named supplies.
Here, then, we have sundry groups of facts directly supporting the belief
that functionally-wrought modifications descend from parents to offspring.
* * * * *
Now let us consider the position of those Darwinians who dissent from
Darwin, and who make light of all this evidence. We might naturally suppose
that their own hypothesis is unassailable. Yet, strange to say, they admit
that there is no direct proof that any species has been established by
natural selection. The proof is inferential only.
The certainty of an axiom does not give certainty to the deductions drawn
from it. That natural selection is, and always has been, operative is
incontestable. Obviously I should be the last person to deny that survival
of the fittest is a necessity: its negation is inconceivable. The
Neo-Darwinians, however, judging from their attitude, apparently assume
that firmness of the basis implies firmness of the superstructure. But
however high may be the probability of some of the conclusions drawn, none
of them can have more than probability; while some of them remain, and are
likely to remain, very questionable. Observe the difficulties.
(1) The general argument proceeds upon the analogy between natural
selection and artificial selection. Yet all know that the first cannot do
what the last does. Natural selection can do nothing more than preserve
those of which the _aggregate_ characters are most favourable to life. It
cannot pick out those possessed of one particular favourable character,
unless this is of extreme importance.
(2) In many cases a structure is of no service until it has reached a
certain development; and it remains to account for that increase of it by
natural selection which must be supposed to take place before it reaches
the stage of usefulness.
(3) Advantageous variations, not preserved in nature as they are by the
breeder, are liable to be swamped by crossing or to disappear by atavism.
Now whatever replies are made, their component propositions cannot be
necessary truths. So that the conclusion in each case, however reasonable,
cannot claim certainty: the fabric can have no stability like that of its
foundation.
When to uncertainties in the arguments supporting the hypothesis we add its
inability to explain facts of cardinal significance, as proved above, there
is I think ground for asserting that natural selection is less clearly
shown to be a factor in the origination of species than is the inheritance
of functionally-wrought changes.
* * * * *
If, finally, it is said that the mode in which functionally-wrought
changes, especially in small parts, so affect the reproductive elements as
to repeat themselves in offspring, cannot be imagined--if it be held
inconceivable that those minute changes in the organs of vision which cause
myopia can be transmitted through the appropriately-modified sperm-cells or
germ-cells; then the reply is that the opposed hypothesis presents a
corresponding inconceivability. Grant that the habit of a pointer was
produced by selection of those in which an appropriate variation in the
nervous system had occurred; it is impossible to imagine how a
slightly-different arrangement of a few nerve-cells and fibres could be
conveyed by a spermatozoon. So too it is impossible to imagine how in a
spermatozoon there can be conveyed the 480,000 independent variables
required for the construction of a single peacock's feather, each having a
proclivity towards its proper place. Clearly the ultimate process by which
inheritance is effected in either case passes comprehension; and in this
respect neither hypothesis has an advantage over the other.
APPENDIX D.
ON ALLEGED "SPONTANEOUS GENERATION," AND ON THE HYPOTHESIS OF PHYSIOLOGICAL
UNITS.
[_The following letter, originally written for publication in the_ North
American Review, _but declined by the Editor in pursuance of a general
rule, and eventually otherwise published in the United States, I have
thought well to append to this first volume of the_ Principles of Biology.
_I do this because the questions which it discusses are dealt with in this
volume; and because the further explanations it furnishes seem needful to
prevent misapprehensions._]
_The Editor of the North American Review._
SIR,
It is in most cases unwise to notice adverse criticisms. Either they do not
admit of answers or the answers may be left to the penetration of readers.
When, however, a critic's allegations touch the fundamental propositions of
a book, and especially when they appear in a periodical having the position
of the _North American Review_, the case is altered. For these reasons the
article on "Philosophical Biology," published in your last number, demands
from me an attention which ordinary criticisms do not.
It is the more needful for me to notice it, because its two leading
objections have the one an actual fairness and the other an apparent
fairness; and in the absence of explanations from me, they will be
considered as substantiated even by many, or perhaps most, of those who
have read the work itself--much more by those who have not read it. That to
prevent the spread of misapprehensions I ought to say something, is further
shown by the fact that the same two objections have already been made in
England--the one by Dr. Child, of Oxford, in his _Essays on Physiological
Subjects_, and the other by a writer in the _Westminster Review_ for July,
1865.
* * * * *
In the note to which your reviewer refers, I have, as he says, tacitly
repudiated the belief in "spontaneous generation;" and that I have done
this in such a way as to leave open the door for the interpretation given
by him is true. Indeed the fact that Dr. Child, whose criticism is a
sympathetic one, puts the same construction on this note, proves that your
reviewer has but drawn what seems to be a necessary inference.
Nevertheless, the inference is one which I did not intend to be drawn.
In explanation, let me at the outset remark that I am placed at a
disadvantage in having had to omit that part of the System of Philosophy
which deals with Inorganic Evolution. In the original programme will be
found a parenthetic reference to this omitted part, which should, as there
stated, precede the _Principles of Biology_. Two volumes are missing. The
closing chapter of the second, were it written, would deal with the
evolution of organic matter--the step preceding the evolution of living
forms. Habitually carrying with me in thought the contents of this
unwritten chapter, I have, in some cases, expressed myself as though the
reader had it before him; and have thus rendered some of my statements
liable to misconstructions. Apart from this, however, the explanation of
the apparent inconsistency is very simple, if not very obvious. In the
first place, I do not believe in the "spontaneous generation" commonly
alleged, and referred to in the note; and so little have I associated in
thought this alleged "spontaneous generation" which I disbelieve, with the
generation by evolution which I do believe, that the repudiation of the one
never occurred to me as liable to be taken for repudiation of the other.
That creatures having _quite specific structures_ are evolved in the course
of a few hours, without antecedents calculated to determine their specific
forms, is to me incredible. Not only the established truths of Biology, but
the established truths of science in general, negative the supposition that
organisms having structures definite enough to identify them as belonging
to known genera and species, can be produced in the absence of germs
derived from antecedent organisms of the same genera and species. If there
can suddenly be imposed on simple protoplasm the organization which
constitutes it a _Paramoecium_, I see no reason why animals of greater
complexity, or indeed of any complexity, may not be constituted after the
same manner. In brief, I do not accept these alleged facts as exemplifying
Evolution, because they imply something immensely beyond that which
Evolution, as I understand it, can achieve. In the second place, my
disbelief extends not only to the alleged cases of "spontaneous
generation," but to every case akin to them. The very conception of
spontaneity is wholly incongruous with the conception of Evolution. For
this reason I regard as objectionable Mr. Darwin's phrase "spontaneous
variation" (as indeed he does himself); and I have sought to show that
there are always assignable causes of variation. No form of Evolution,
inorganic or organic, can be spontaneous; but in every instance the
antecedent forces must be adequate in their quantities, kinds, and
distributions, to work the observed effects. Neither the alleged cases of
"spontaneous generation," nor any imaginable cases in the least allied to
them, fulfil this requirement.
If, accepting these alleged cases of "spontaneous generation," I had
assumed, as your reviewer seems to do, that the evolution of organic life
commenced in an analogous way; then, indeed, I should have left myself open
to a fatal criticism. This supposed "spontaneous generation" habitually
occurs in menstrua that contain either organic matter, or matter originally
derived from organisms; and such organic matter, proceeding in all known
cases from organisms of a higher kind, implies the pre-existence of such
higher organisms. By what kind of logic, then, is it inferrible that
organic life was initiated after a manner like that in which _Infusoria_
are said to be now spontaneously generated? Where, before life commenced,
were the superior organisms from which these lowest organisms obtained
their organic matter? Without doubting that there are those who, as the
reviewer says, "can penetrate deeper than Mr. Spencer has done into the
idea of universal evolution," and who, as he contends, prove this by
accepting the doctrine of "spontaneous generation"; I nevertheless think
that I can penetrate deep enough to see that a tenable hypothesis
respecting the origin of organic life must be reached by some other clue
than that furnished by experiments on decoction of hay and extract of beef.
From what I do not believe, let me now pass to what I do believe. Granting
that the formation of organic matter, and the evolution of life in its
lowest forms, may go on under existing cosmical conditions; but believing
it more likely that the formation of such matter and such forms, took place
at a time when the heat of the Earth's surface was falling through those
ranges of temperature at which the higher organic compounds are unstable; I
conceive that the moulding of such organic matter into the simplest types,
must have commenced with portions of protoplasm more minute, more
indefinite, and more inconstant in their characters, than the lowest
Rhizopods--less distinguishable from a mere fragment of albumen than even
the _Protogenes_ of Professor Haeckel. The evolution of specific shapes
must, like all other organic evolution, have resulted from the actions and
reactions between such incipient types and their environments, and the
continued survival of those which happened to have specialities best fitted
to the specialities of their environments. To reach by this process the
comparatively well-specialized forms of ordinary _Infusoria_, must, I
conceive, have taken an enormous period of time.
To prevent, as far as may be, future misapprehension, let me elaborate this
conception so as to meet the particular objections raised. The reviewer
takes for granted that a "first organism" must be assumed by me, as it is
by himself. But the conception of a "first organism," in anything like the
current sense of the words, is wholly at variance with conception of
evolution; and scarcely less at variance with the facts revealed by the
microscope. The lowest living things are not properly speaking organisms at
all; for they have no distinctions of parts--no traces of organization. It
is almost a misuse of language to call them "forms" of life: not only are
their outlines, when distinguishable, too unspecific for description, but
they change from moment to moment and are never twice alike, either in two
individuals or in the same individual. Even the word "type" is applicable
in but a loose way; for there is little constancy in their generic
characters: according as the surrounding conditions determine, they undergo
transformations now of one kind and now of another. And the vagueness, the
inconstancy, the want of appreciable structure, displayed by the simplest
of living things as we now see them, are characters (or absences of
characters) which, on the hypothesis of Evolution, must have been still
more decided when, as at first, no "forms," no "types," no "specific
shapes," had been moulded. That "absolute commencement of organic life on
the globe," which the reviewer says I "cannot evade the admission of," I
distinctly deny. The affirmation of universal evolution is in itself the
negation of an "absolute commencement" of anything. Construed in terms of
evolution, every kind of being is conceived as a product of modifications
wrought by insensible gradations on a pre-existing kind of being; and this
holds as fully of the supposed "commencement of organic life" as of all
subsequent developments of organic life. It is no more needful to suppose
an "absolute commencement of organic life" or a "first organism," than it
is needful to suppose an absolute commencement of social life and a first
social organism. The assumption of such a necessity in this last case, made
by early speculators with their theories of "social contracts" and the
like, is disproved by the facts; and the facts, so far as they are
ascertained, disprove the assumption of such a necessity in the first case.
That organic matter was not produced all at once, but was reached through
steps, we are well warranted in believing by the experiences of chemists.
Organic matters are produced in the laboratory by what we may literally
call _artificial evolution_. Chemists find themselves unable to form these
complex combinations directly from their elements; but they succeed in
forming them indirectly, by successive modifications of simpler
combinations. In some binary compound, one element of which is present in
several equivalents, a change is made by substituting for one of these
equivalents an equivalent of some other element; so producing a ternary
compound. Then another of the equivalents is replaced, and so on. For
instance, beginning with ammonia, N H_{3}, a higher form is obtained by
replacing one of the atoms of hydrogen by an atom of methyl, so producing
methyl-amine, N (C H_{3} H_{2}); and then, under the further action of
methyl, ending in a further substitution, there is reached the still more
compound substance dimethyl-amine, N (C H_{3}) (C H_{3}) H. And in this
manner highly complex substances are eventually built up. Another
characteristic of their method is no less significant. Two complex
compounds are employed to generate, by their action upon one another, a
compound of still greater complexity: different heterogeneous molecules of
one stage, become parents of a molecule a stage higher in heterogeneity.
Thus, having built up acetic acid out of its elements, and having by the
process of substitution described above, changed the acetic acid into
propionic acid, and propionic into butyric, of which the formula is
{C(CH_{3})(CH_{3})H}
{CO(HO) };
this complex compound, by operating on another complex compound, such as
the dimethyl-amine named above, generates one of still greater complexity,
butyrate of dimethyl-amine
{C(CH)(CH_{3})H} N(CH_{3})(CH_{3})H.
{CO(HO) }
See, then, the remarkable parallelism. The progress towards higher types of
organic molecules is effected by modifications upon modifications; as
throughout Evolution in general. Each of these modifications is a change of
the molecule into equilibrium with its environment--an adaptation, as it
were, to new surrounding conditions to which it is subjected; as throughout
Evolution in general. Larger, or more integrated, aggregates (for compound
molecules are such) are successively generated; as throughout Evolution in
general. More complex or heterogeneous aggregates are so made to arise, one
out of another; as throughout Evolution in general. A
geometrically-increasing multitude of these larger and more complex
aggregates so produced, at the same time results; as throughout Evolution
in general. And it is by the action of the successively higher forms on one
another, joined with the action of environing conditions, that the highest
forms are reached; as throughout Evolution in general.
When we thus see the identity of method at the two extremes--when we see
that the general laws of evolution, as they are exemplified in known
organisms, have been unconsciously conformed to by chemists in the
artificial evolution of organic matter; we can scarcely doubt that these
laws were conformed to in the natural evolution of organic matter, and
afterwards in the evolution of the simplest organic forms. In the early
world, as in the modern laboratory, inferior types of organic substances,
by their mutual actions under fit conditions, evolved the superior types of
organic substances, ending in organizable protoplasm. And it can hardly be
doubted that the shaping of organizable protoplasm, which is a substance
modifiable in multitudinous ways with extreme facility, went on after the
same manner. As I learn from one of our first chemists, Prof. Frankland,
_protein_ is capable of existing under probably at least a thousand
isomeric forms; and, as we shall presently see, it is capable of forming,
with itself and other elements, substances yet more intricate in
composition, that are practically infinite in their varieties of kind.
Exposed to those innumerable modifications of conditions which the Earth's
surface afforded, here in amount of light, there in amount of heat, and
elsewhere in the mineral quality of its aqueous medium, this extremely
changeable substance must have undergone now one, now another, of its
countless metamorphoses. And to the mutual influences of its metamorphic
forms under favouring conditions, we may ascribe the production of the
still more composite, still more sensitive, still more variously-changeable
portions of organic matter, which, in masses more minute and simpler than
existing _Protozoa_, displayed actions verging little by little into those
called vital--actions which protein itself exhibits in a certain degree,
and which the lowest known living things exhibit only in a greater degree.
Thus, setting out with inductions from the experiences of organic chemists
at the one extreme, and with inductions from the observations of biologists
at the other extreme, we are enabled deductively to bridge the
interval--are enabled to conceive how organic compounds were evolved, and
how, by a continuance of the process, the nascent life displayed in these
became gradually more pronounced. And this it is which has to be explained,
and which the alleged cases of "spontaneous generation" would not, were
they substantiated, help us in the least to explain.
It is thus manifest, I think, that I have not fallen into the alleged
inconsistency. Nevertheless, I admit that your reviewer was justified in
inferring this inconsistency; and I take blame to myself for not having
seen that the statement, as I have left it, is open to misconstruction.
* * * * *
I pass now to the second allegation--that in ascribing to certain specific
molecules, which I have called "physiological units," the aptitude to build
themselves into the structure of the organism to which they are peculiar, I
have abandoned my own principle, and have assumed something beyond the
re-distribution of Matter and Motion. As put by the reviewer, his case
appears to be well made out; and that he is not altogether unwarranted in
so putting it, may be admitted. Nevertheless, there does not in reality
exist the supposed incongruity.
Before attempting to make clear the adequacy of the conception which I am
said to have tacitly abandoned as insufficient, let me remove that excess
of improbability the reviewer gives to it, by the extremely-restricted
meaning with which he uses the word mechanical. In discussing a proposition
of mine he says:--
"He then cites certain remarks of Mr. Paget on the permanent effects
wrought in the blood by the poison of scarlatina and small-pox, as
justifying the belief that such a 'power' exists, and attributes the
repair of a wasted tissue to 'forces analogous to those by which a
crystal reproduces its lost apex.' (Neither of which phenomena, however,
is explicable by mechanical causes.)"
Were it not for the deliberation with which this last statement is made, I
should take it for a slip of the pen. As it is, however, I have no course
left but to suppose the reviewer unaware of the fact that molecular actions
of all kinds are now not only conceived as mechanical actions, but that
calculations based on this conception of them, bring out the results that
correspond with observation. There is no kind of re-arrangement among
molecules (crystallization being one) which the modern physicist does not
think of. and correctly reason upon, in terms of forces and motions like
those of sensible masses. Polarity is regarded as a resultant of such
forces and motions; and when, as happens in many cases, light changes the
molecular structure of a crystal, and alters its polarity, it does this by
impressing, in conformity with mechanical laws, new motions on the
constituent molecules. That the reviewer should present the mechanical
conception under so extremely limited a form, is the more surprising to me
because, at the outset of the very work he reviews, I have, in various
passages, based inferences on those immense extensions of it which he
ignores; indicating, for example, the interpretation it yields of the
inorganic chemical changes effected by heat, and the organic chemical
changes effected by light (_Principles of Biology_, § 13).
Premising, then, that the ordinary idea of mechanical action must be
greatly expanded, let us enter upon the question at issue--the sufficiency
of the hypothesis that the structure of each organism is determined by the
polarities of the special molecules, or physiological units, peculiar to it
as a species, which necessitate tendencies towards special arrangements. My
proposition and the reviewer's criticism upon it, will be most conveniently
presented if I quote in full a passage of his from which I have already
extracted some expressions. He says:--
"It will be noticed, however, that Mr. Spencer attributes the possession
of these 'tendencies,' or 'proclivities,' to natural inheritance from
ancestral organisms; and it may be argued that he thus saves the
mechanist theory and his own consistency at the same time, inasmuch as he
derives even the 'tendencies' themselves ultimately from the environment.
To this we reply, that Mr. Spencer, who advocates the nebular hypothesis,
cannot evade the admission of an absolute commencement of organic life on
the globe, and that the 'formative tendencies,' without which he cannot
explain the evolution of a single individual, could not have been
inherited by the first organism. Besides, by his virtual denial of
spontaneous generation, he denies that the first organism was evolved out
of the inorganic world, and thus shuts himself off from the argument
(otherwise plausible) that its 'tendencies' were ultimately derived from
the environment."
This assertion is already in great measure disposed of by what has been
said above. Holding that, though not "spontaneously generated," those
minute portions of protoplasm which first displayed in the feeblest degree
that changeability taken to imply life, were evolved, I am _not_ debarred
from the argument that the "tendencies" of the physiological units are
derived from the inherited effects of environing actions. If the conception
of a "first organism" were a necessary one, the reviewer's objection would
be valid. If there were an "absolute commencement" of life, a definite line
parting organic matter from the simplest living forms, I should be placed
in the predicament he describes. But as the doctrine of Evolution itself
tacitly negatives any such distinct separation; and as the negation is the
more confirmed by the facts the more we know of them; I do not feel that I
am entangled in the alleged difficulty. My reply might end here; but as the
hypothesis in question is one not easily conceived, and very apt to be
misunderstood, I will attempt a further elucidation of it.
Much evidence now conspires to show that molecules of the substances we
call elementary are in reality compound; and that, by the combination of
these with one another, and re-combinations of the products, there are
formed systems of systems of molecules, unimaginable in their complexity.
Step by step as the aggregate molecules so resulting, grow larger and
increase in heterogeneity, they become more unstable, more readily
transformable by small forces, more capable of assuming various characters.
Those composing organic matter transcend all others in size and intricacy
of structure; and in them these resulting traits reach their extreme. As
implied by its name _protein_, the essential substance of which organisms
are built, is remarkable alike for the variety of its metamorphoses and the
facility with which it undergoes them: it changes from one to another of
its thousand isomeric forms on the slightest change of conditions. Now
there are facts warranting the belief that though these multitudinous
isomeric forms of protein will not unite directly with one another, yet
they admit of being linked together by other elements with which they
combine. And it is very significant that there are habitually present two
other elements, sulphur and phosphorus, which have quite special powers of
holding together many equivalents--the one being pentatomic and the other
hexatomic. So that it is a legitimate supposition (justified by analogies)
that an atom of sulphur may be a bond of union among half-a-dozen different
isomeric forms of protein; and similarly with phosphorus. A moment's
thought will show that, setting out with the thousand isomeric forms of
protein, this makes possible a number of these combinations almost passing
the power of figures to express. Molecules so produced, perhaps exceeding
in size and complexity those of protein as those of protein exceed those of
inorganic matter, may, I conceive, be the special units belonging to
special kinds of organisms. By their constitution they must have a
plasticity, or sensitiveness to modifying forces, far beyond that of
protein; and bearing in mind not only that their varieties are practically
infinite in number, but that closely allied forms of them, chemically
indifferent to one another as they must be, may coexist in the same
aggregate, we shall see that they are fitted for entering into unlimited
varieties of organic structures.
The existence of such physiological units, peculiar to each species of
organism, is not unaccounted for. They are evolved simultaneously with the
evolution of the organisms they compose--they differentiate as fast as
these organisms differentiate; and are made multitudinous in kind by the
same actions which make the organism they compose multitudinous, in kind.
This conception is clearly representable in terms of the mechanical
hypothesis. Every physicist will endorse the proposition that in each
aggregate there tends to establish itself an equilibrium between the forces
exercised by all the units upon each and by each upon all. Even in masses
of substance so rigid as iron and glass, there goes on a molecular
re-arrangement, slow or rapid according as circumstances facilitate, which
ends only when there is a complete balance between the actions of the parts
on the whole and the actions of the whole on the parts: the implications
being that every change in the form or size of the whole, necessitates some
redistribution of the parts. And though in cases like these, there occurs
only a polar re-arrangement of the molecules, without changes in the
molecules themselves; yet where, as often happens, there is a passage from
the colloid to the crystalloid state, a change of constitution occurs in
the molecules themselves. These truths are not limited to inorganic matter:
they unquestionably hold of organic matter. As certainly as molecules of
alum have a form of equilibrium, the octahedron, into which they fall when
the temperature of their solvent allows them to aggregate, so certainly
must organic molecules of each kind, no matter how complex, have a form of
equilibrium in which, when they aggregate, their complex forces are
balanced--a form far less rigid and definite, for the reason that they have
far less definite polarities, are far more unstable, and have their
tendencies more easily modified by environing conditions. Equally certain
is it that the special molecules having a special organic structure as
their form of equilibrium, must be reacted upon by the total forces of this
organic structure; and that, if environing actions lead to any change in
this organic structure, these special molecules, or physiological units,
subject to a changed distribution of the total forces acting upon them will
undergo modification--modification which their extreme plasticity will
render easy. By this action and reaction I conceive the physiological units
peculiar to each kind of organism, to have been moulded along with the
organism itself. Setting out with the stage in which protein in minute
aggregates, took on those simplest differentiations which fitted it for
differently-conditioned parts of its medium, there must have unceasingly
gone on perpetual re-adjustments of balance between aggregates and their
units--actions and reactions of the two, in which the units tended ever to
establish the typical form produced by actions and reactions in all
antecedent generations, while the aggregate, if changed in form by change
of surrounding conditions, tended ever to impress on the units a
corresponding change of polarity, causing them in the next generation to
reproduce the changed form--their new form of equilibrium.
This is the conception which I have sought to convey, though it seems
unsuccessfully, in the _Principles of Biology_; and which I have there used
to interpret the many involved and mysterious phenomena of Genesis,
Heredity, and Variation. In one respect only am I conscious of having so
inadequately explained myself, as to give occasion for a
misinterpretation--the one made by the _Westminster_ reviewer above
referred to. By him, as by your own critic, it is alleged that in the idea
of "inherent tendencies" I have introduced, under a disguise, the
conception of "the archæus, vital principle, _nisus formativus_, and so
on." This allegation is in part answered by the foregoing explanation. That
which I have here to add, and did not adequately explain in the _Principles
of Biology_, is that the proclivity of units of each order towards the
specific arrangement seen in the organism they form, is not to be
understood as resulting from their own structures and actions only; but as
the product of these and the environing forces to which they are exposed.
Organic evolution takes place only on condition that the masses of
protoplasm formed of the physiological units, and of the assimilable
materials out of which others like themselves are to be multiplied, are
subject to heat of a given degree--are subject, that is, to the unceasing
impacts of undulations of a certain strength and period; and, within
limits, the rapidity with which the physiological units pass from their
indefinite arrangement to the definite arrangement they presently assume,
is proportionate to the strengths of the ethereal undulations falling upon
them. In its complete form, then, the conception is that these specific
molecules, having the immense complexity above described, and having
correspondently complex polarities which cannot be mutually balanced by any
simple form of aggregation, have, for the form of aggregation in which all
their forces are equilibrated, the structure of the adult organism to which
they belong; and that they are compelled to fall into this structure by the
co-operation of the environing forces acting on them, and the forces they
exercise on one another--the environing forces being the source of the
_power_ which effects the re-arrangement, and the polarities of the
molecules determining the _direction_ in which that power is turned. Into
this conception there enters no trace of the hypothesis of an "archæus or
vital principle;" and the principles of molecular physics fully justify it.
It is, however, objected that "the living body in its development presents
a long succession of _differing_ forms; a continued series of changes for
the whole length of which, according to Mr. Spencer's hypothesis, the
physiological units must have an 'inherent tendency.' Could we more truly
say of anything, 'it is unrepresentable in thought?'" I reply that if there
is taken into account an element here overlooked, the process will not be
found "unrepresentable in thought." This is the element of size or mass. To
satisfy or balance the polarities of each order of physiological units, not
only a certain structure of organism, but a certain size of organism is
needed; for the complexities of that adult structure in which the
physiological units are equilibrated, cannot be represented within the
small bulk of the embryo. In many minute organisms, where the whole mass of
physiological units required for the structure is present, the very thing
_does_ take place which it is above implied _ought_ to take place. The mass
builds itself directly into the complete form. This is so with _Acari_, and
among the nematoid _Entozoa_. But among higher animals such direct
transformations cannot happen. The mass of physiological units required to
produce the size as well as the structure that approximately equilibrates
them, is not all present, but has to be formed by successive
additions--additions which in viviparous animals are made by absorbing, and
transforming into these special molecules, the organizable materials
directly supplied by the parent, and which in oviparous animals are made by
doing the like with the organizable materials in the "food-yelk," deposited
by the parent in the same envelope with the germ. Hence it results that,
under such conditions, the physiological units which first aggregate into
the rudiment of the future organism, do not form a structure like that of
the adult organism, which, when of such small dimensions, does not
equilibrate them. They distribute themselves so as partly to satisfy the
chief among their complex polarities. The vaguely-differentiated mass thus
produced cannot, however, be in equilibrium. Each increment of
physiological units formed and integrated by it, changes the distribution
of forces; and this has a double effect. It tends to modify the
differentiations already made, bringing them a step nearer to the
equilibrating structure; and the physiological units next integrated, being
brought under the aggregate of polar forces exercised by the whole mass,
which now approaches a step nearer to that ultimate distribution of polar
forces which exists in the adult organism, are coerced more directly into
the typical structure. Thus there is necessitated a series of compromises.
Each successive form assumed is unstable and transitional: approach to the
typical structure going on hand in hand with approach to the typical bulk.
Possibly I have not succeeded by this explanation, any more than by the
original explanation, in making this process "representable in thought." It
is manifestly untrue, however, that I have, as alleged, re-introduced under
a disguise the conception of a "vital principle." That I interpret
embryonic development in terms of Matter and Motion, cannot, I think, be
questioned. Whether the interpretation is adequate, must be a matter of
opinion; but it is clearly a matter of fact, that I have not fallen into
the inconsistency asserted by your reviewer. At the same time I willingly
admit that, in the absence of certain statements which I have now supplied,
he was not unwarranted in representing my conception in the way that he has
done.
----
NOTES
[1] Gross misrepresentations of this statement, which have been from time
to time made, oblige me, much against my will, to add here an
explanation of it. The last of these perversions, uttered in a
lecture delivered at Belfast by the Rev. Professor Watts, D.D., is
reported in the _Belfast Witness_ of December 18, 1874; just while a
third impression of this work is being printed from the plates. The
report commences as follows:--"Dr. Watts, after showing that on his
own confession Spencer was indebted for his facts to Huxley and
Hooker, who," &c., &c.
Wishing in this, as in other cases, to acknowledge indebtedness when
conscious of it, I introduced the words referred to, in recognition
of the fact that I had repeatedly questioned the distinguished
specialists named, on matters beyond my knowledge, which were not
dealt with in the books at my command. Forgetting the habits of
antagonists, and especially theological antagonists, it never
occurred to me that my expression of thanks to my friends for
"information where my own was deficient," would be turned into the
sweeping statement that I was indebted to them for my facts.
Had Professor Watts looked at the preface to the second volume (the
two having been published separately, as the prefaces imply), he
would have seen a second expression of my indebtedness "for their
valuable criticisms, and for the trouble they have taken in
_checking_ the numerous statements of fact on which the arguments
proceed"--no further indebtedness being named. A moment's comparison
of the two volumes in respect of their accumulations of facts, would
have shown him what kind of warrant there was for his interpretation.
Doubtless the Rev. Professor was prompted to make this assertion by
the desire to discredit the work he was attacking; and having so good
an end in view, thought it needless to be particular about the means.
In the art of dealing with the language of opponents, Dr. Watts might
give lessons to Monsignor Capel and Archbishop Manning.
_December 28th, 1874._
[2] In this passage as originally written (in 1862) they were described
as incondensible; since, though reduced to the density of liquids,
they had not been liquefied.
[3] Here and hereafter the word "atom" signifies a unit of something
classed as an element, because thus far undecomposed by us. The word
must not be supposed to mean that which its derivation implies. In
all probability it is not a simple unit but a compound one.
[4] The name hydro-carbons was here used when these pages were written,
thirty-four years ago. It was the name then current. In this case, as
in multitudinous other cases, the substitution of newer words and
phrases for older ones, is somewhat misleading. Putting the thoughts
of 1862 in the language of 1897 gives an illusive impression of
recency.
[5] It will perhaps seem strange to class oxygen as a crystalloid. But
inasmuch as the crystalloids are distinguished from the colloids by
their atomic simplicity, and inasmuch as sundry gases are reducible
to a crystalline state, we are justified in so classing it.
[6] The remark made by a critic to the effect that in a mammal higher
temperature diminishes the rate of molecular change in the tissues,
leads me to add that the exhalation I have alleged is prevented if
the heat rises above the range of variation normal to the organism;
since, then, unusually rapid pulsations with consequent inefficient
propulsion of the blood, cause a diminished rate of circulation. To
produce the effect referred to in the text, heat must be associated
with dryness; for otherwise evaporation is not aided. General
evidence supporting the statement I have made is furnished by the
fact that the hot and dry air of the eastern deserts is extremely
invigorating; by the fact that all the energetic and conquering races
of men have come from the hot and dry regions marked on the maps as
rainless; and by the fact that travellers in Africa comment on the
contrast between the inhabitants of the hot and dry regions
(relatively elevated) and those of the hot and moist regions: active
and inert respectively.
[7] The increase of respiration found to result from the presence of
light, is probably an _indirect_ effect. It is most likely due to the
reception of more vivid impressions through the eyes, and to the
consequent nervous stimulation. Bright light is associated in our
experience with many of our greatest outdoor pleasures, and its
presence partially arouses the consciousness of them, with the
concomitant raised vital functions.
[8] To exclude confusion it may be well here to say that the word "atom"
is, as before explained, used as the name for a unit of a substance
at present undecomposed; while the word "molecule" is used as the
name for a unit of a substance known to be compound.
[9] On now returning to the subject after many years, I meet with some
evidence recently assigned, in a paper read before the Royal Society
by Mr. J. W. Pickering, D.Sc. (detailing results harmonizing with
those obtained by Prof. Grimaux), showing clearly how important an
agent in vital actions is this production of isomeric changes by
slight changes of conditions. Certain artificially produced
substances, simulating proteids in other of their characters and
reactions, were found to simulate them in coagulability by trifling
disturbances. "In the presence of a _trace of neutral salt_ they
coagulate on heating at temperatures very similar to proteid
solutions." And it is shown that by one of these factitious organic
colloids a like effect is produced in coagulating the blood, to that
"produced by the intravenous injection of a nucleoproteid."
[10] After this long interval during which other subjects have occupied
me, I now find that the current view is similar to the view above set
forth, in so far that a small molecular disturbance is supposed
suddenly to initiate a great one, producing a change compared to an
explosion. But while, of two proposed interpretations, one is that
the fuse is nitrogenous and the charge a carbo-hydrate, the other is
that both are nitrogenous. The relative probabilities of these
alternative views will be considered in a subsequent chapter.
[11] When writing this passage I omitted to observe the verification
yielded of the conclusion contained in § 15 concerning the part
played in the vital processes by the nitrogenous compounds. For these
vegeto-alkalies, minute quantities of which produce such great
effects in exalting the functions (_e. g._, a sixteenth of a grain of
strychnia is a dose), are all nitrogenous bodies, and, by
implication, relatively unstable bodies. The small amounts of
molecular change which take place in these small quantities of the
vegeto-alkalies when diffused through the system, initiate larger
amounts of molecular change in the nitrogenous elements of the
tissues.
But the evidence furnished a generation ago by these vegeto-alkalies
has been greatly reinforced by far more striking evidence furnished
by other nitrogenous compounds--the various explosives. These, at the
same time that they produce by their sudden decompositions violent
effects outside the organism, also produce violent effects inside it:
a hundredth of a grain of nitro-glycerine being a sufficient dose.
Investigations made by Dr. J. B. Bradbury, and described by him in
the Bradshaw Lecture on "Some New Vaso-Dilators" (see _The Lancet_,
Nov. 16, 1895), details the effects of kindred
bodies--methyl-nitrate, glycol-dinitrate, erythrol-tetranitrate. The
first two, in common with nitro-glycerine, are stable only when cool
and in the dark--sunlight or warmth decomposes them, and they explode
by rapid heating or percussion. The fact which concerns us here is
that the least stable--glycol-dinitrate--has the most powerful and
rapid physiological effect, which is proportionately transient. In
one minute the blood-pressure is reduced by one-fourth and in four
minutes by nearly two-thirds: an effect which is dissipated in a
quarter of an hour. So that this excessively unstable compound,
decomposing in the body in a very short time, produces within that
short time a vast amount of molecular change: acting, as it seems,
not through the nervous system, but directly on the blood-vessels.
[12] This interpretation is said to be disproved by the fact that the
carbo-hydrate contained in muscle amounts to only about 1.5 of the
total solids. I do not see how this statement is to be reconciled
with the statement cited three pages back from Professor Michael
Foster, that the deposits of glycogen contained in the liver and in
the muscles may be compared to the deposits in a central bank and
branch banks.
[13] Before leaving the topic let me remark that the doctrine of
metabolism is at present in its inchoate stage, and that the
prevailing conclusions should be held tentatively. As showing this
need an anomalous fact may be named. It was long held that gelatine
is of small value as food, and though it is now recognized as
valuable because serving the same purposes as fats and
carbo-hydrates, it is still held to be valueless for structural
purposes (save for some inactive tissue); and this estimate agrees
with the fact that it is a relatively stable nitrogenous compound,
and therefore unfit for those functions performed by unstable
nitrogenous compounds in the muscular and other tissues. But if this
is true, it seems a necessary implication that such substances as
hair, wool, feathers, and all dermal growths chemically akin to
gelatine, and even more stable, ought to be equally innutritive or
more innutritive. In that case, however, what are we to say of the
larva of the clothes-moth, which subsists exclusively on one or other
of these substances, and out of it forms all those unstable
nitrogenous compounds needful for carrying on its life and developing
its tissues? Or again, how are we to understand the nutrition of the
book-worm, which, in the time-stained leaves through which it
burrows, finds no proteid save that contained in the dried-up size,
which is a form of gelatine; or, once more, in what form is the
requisite amount of nitrogenous substance obtained by the
coleopterous larva which eats holes in wood a century old?
[14] This chapter and the following two chapters originally appeared in
Part III of the original edition of the _Principles of Psychology_
(1855): forming a preliminary which, though indispensable to the
argument there developed, was somewhat parenthetical. Having now to
deal with the general science of Biology before the more special one
of Psychology, it becomes possible to transfer these chapters to
their proper place.
[15] See _Westminster Review_ for April, 1852.--Art. IV. "A Theory of
Population." See Appendix A.
[16] This paragraph replaces a sentence that, in _The Principles of
Psychology_, referred to a preceding chapter on "Method;" in which
the mode of procedure here indicated was set forth as a mode to be
systematically pursued in the choice of hypotheses. This chapter on
Method is now included, along with other matter, in a volume entitled
_Various Fragments_.
[17] Speaking of "the general idea of _life_" M. Comte says:--"Cette idée
suppose, en effet, non-seulement celle d'un être organisé de manière
à comporter l'état vital, mais aussi celle, non moins indispensable,
d'un certain ensemble d'influences extérieures propres à son
accomplissement. Une telle harmonie entre l'être vivant et le
_milieu_ correspondant, caractérise evidemment la condition
fondamentale de la vie." Commenting on de Blainville's definition of
life, which he adopts, he says:--"Cette lumineuse définition ne me
paraît laisser rien d'important à désirer, si ce n'est une indication
plus directe et plus explicite de ces deux conditions fondamentales
co-relatives, nécessairement inséparables de l'état vivant, un
_organisme_ déterminé et un _milieu_ convenable." It is strange that
M. Comte should have thus recognized the necessity of a harmony
between an organism and its environment, as a _condition_ essential
to life, and should not have seen that the continuous maintenance of
such inner actions as will counterbalance outer actions,
_constitutes_ life.
[When the original edition was published Dr. J. H. Bridges wrote to
me saying that in the _Politique Positive_, Comte had developed his
conception further. On p. 413, denying "le prétendu antagonisme des
corps vivants envers leurs milieux inorganiques," he says "au lieu de
ce conflit, on a reconnu bientôt que cette relation nécessaire
constitue une condition fondamentale de la vie réelle, dont la notion
systématique consiste dans une intime conciliation permanente entre
la spontanéité intérieure et la fatalité extérieure." Still, this
"conciliation _permanente_" seems to be a "_condition_" to life; not
that varying adjustment of changes which life consists in
maintaining. In presence of an ambiguity, the interpretation which
agrees with his previous statement must be chosen.]
[18] In further elucidation of this general doctrine, see _First
Principles_, § 25.
[19] In ordinary speech Development is often used as synonymous with
Growth. It hence seems needful to say that Development as here and
hereafter used, means _increase of structure_ and not _increase of
bulk_. It may be added that the word Evolution, comprehending growth
as well as Development, is to be reserved for occasions when both are
implied.
[20] This paragraph originally formed part of a review-article on
"Transcendental Physiology," published in 1857.
[21] When, in 1863, the preceding chapter was written, it had not occurred
to me that there needed an accompanying chapter treating of
Structure. The gap left by that oversight I now fill up. In doing
this there have been included certain statements which are tacitly
presupposed in the last chapter, and there may also be some which
overlap statements in the next chapter. I have not thought it needful
so to alter adjacent chapters as to remove these slight defects: the
duplicated ideas will bear re-emphasizing.
[22] In connexion with this matter I add here a statement made by Prof.
Foster which it is difficult to understand: "Indeed it has been
observed that a dormouse actually gained in weight during a
hybernating period; it discharged during this period neither urine
nor fæces, and the gain in weight was the excess of oxygen taken in
over the carbonic acid given out." (_Text-book of Physiology_, 6th
ed., Part II, page 859.)
[23] In the account of James Mitchell, a boy born blind and deaf, given by
James Wardrop, F.R.S. (Edin. 1813), it is said that he acquired a
"preternatural acuteness of touch and smell." The deaf Dr. Kitto
described himself as having an extremely strong visual memory: he
retained "a clear impression or image of everything at which he ever
looked."
[24] Here, as in sundry places throughout this chapter, the necessities of
the argument have obliged me to forestall myself, by assuming the
conclusion reached in a subsequent chapter, that modifications of
structure produced by modifications of function are transmitted to
offspring.
[25] Whether the _Volvox_ is to be classed as animal or vegetal is a
matter of dispute; but its similarity to the blastula stage of many
animals warrants the claim of the zoologists.
[26] While the proof was in my hands there was published in _Science
Progress_ an essay by Dr. T. G. Brodie on "The Phosphorus-containing
Substances of the Cell." In this essay it is pointed out that
"nucleic acid is particularly characterized by its instability.... In
the process of purification it is extremely liable to decompose, with
the result that it loses a considerable part of its phosphorus. In
the second place it is most easily split up in another manner in
which it loses a considerable part of its nitrogen.... To avoid the
latter source of error he [Miescher] found that it was necessary to
keep the temperature of all solutions down to 0°C., the whole time of
the preparation." These facts tend strongly to verify the hypothesis
that the nucleus is a source of perpetual molecular disturbance--not
a regulating centre but a stimulating centre.
[27] The writing of the above section reminded me of certain allied views
which I ventured to suggest nearly 50 years ago. They are contained
in the _Westminster Review_ for April, 1852, in an article entitled
"A Theory of Population deduced from the General Law of Animal
Fertility." It is there suggested that the "spermatozoon is
essentially a neural element, and the ovum essentially a hæmal
element," or, as otherwise stated, that the "sperm-cell is
co-ordinating matter and the germ-cell matter to be co-ordinated"
(pp. 490-493). And along with this proposition there is given some
chemical evidence tending to support it. Now if, in place of "neural"
and "hæmal," we say--the element that is most highly phosphorized and
the element that is phosphorized in a much smaller degree; or if, in
place of co-ordinating matter and matter to be co-ordinated, we
say--the matter which initiates action and the matter which is made
to act; there is disclosed a kinship between this early view and the
view just set forth. In the last part of this work, "Laws of
Multiplication," which is developed from the essay referred to, I
left out the portion containing the quoted sentences, and the
evidence supporting the conclusion drawn. Partly I omitted them
because the speculation did not form an essential link in the general
argument, and partly because I did not see how the suggested
interpretation could hold of plants as well as of animals. If,
however, the alleged greater staining capacity of the male generative
nucleus in plants implies, as in other cases, that the male cell has
a larger proportion of the phosphorized matter than the other
elements concerned, then the difficulty disappears.
As, along with the idea just named, the dropped portion of the
original essay contains other ideas which seem to me worth
preserving, I have thought it as well to reproduce it, in company
with the chief part of the general argument as at first sketched out.
It will be found in Appendix A to this volume.
[28] Unfortunately the word _heterogenesis_ has been already used as a
synonym for "spontaneous generation." Save by those few who believe
in "spontaneous generation," however, little objection will be felt
to using the word in a sense that seems much more appropriate. The
meaning above given to it covers both Metagenesis and
Parthenogenesis.
[29] Prof. Huxley avoids this difficulty by making every kind of Genesis a
mode of development. His classification, which suggested the one
given above, is as follows:--
{ Growth
{ Continuous {
{ { Metamorphosis
{
Development {
{ { Metagenesis
{ { Agamogenesis {
{ Discontinuous { { Parthenogenesis
{ Gamogenesis
[30] The implication is that an essentially similar process occurs in
those fragments of leaves used for artificial propagation. Besides
the Begonias in general, I learn that various other plants are thus
multiplied--Citron and orange trees, _Hoya carnosa_, _Aucuba
japonica_, _Clianthus puniceus_, etc., etc. _Bryophyllum calicinum_,
_Rochea falcata_, and _Echeveria_. I also learn that the following
plants, among others, produce buds from their foliage
leaves:--_Cardamine pratensis_, _Nasturtium officinale_, _Roripa
palustris_, _Brassica oleracea_, _Arabis pumila_, _Chelidonium
majus_, _Nymphæa guianensis_, _Episcia bicolor_, _Chirita sivensis_,
_Pinguicula Backeri_, _Allium_, _Gagea_, _Tolmia_, _Fritillaria_,
_Ornithogalum_, etc. In _Cardamine_ and several others, a complete
miniature plant is at once produced; in other cases bulbils or
similar detachable buds.
[31] Among various examples I have observed, the most remarkable were
among Foxgloves, growing in great numbers and of large size, in a
wood between Whatstandwell Bridge and Crich, in Derbyshire. In one
case the lowest flower on the stem contained, in place of a pistil, a
shoot or spike of flower-buds, similar in structure to the
embryo-buds of the main spike. I counted seventeen buds on it; of
which the first had three stamens, but was otherwise normal; the
second had three; the third, four; the fourth, four; &c. Another
plant, having more varied monstrosities, evinced excess of nutrition
with equal clearness. The following are the notes I took of its
structure:--1st, or lowest flower on the stem, very large; calyx
containing eight divisions, one partly transformed into a corolla,
and another transformed into a small bud with bract (this bud
consisted of a five-cleft calyx, four sessile anthers, a pistil, and
a rudimentary corolla); the corolla of the main flower, which was
complete, contained six stamens, three of them bearing anthers, two
others being flattened and coloured, and one rudimentary; there was
no pistil but, _in place of it_, a large bud, consisting of a
three-cleft calyx of which two divisions were tinted at the ends, an
imperfect corolla marked internally with the usual purple spots and
hairs, three anthers sessile on this mal-formed corolla, a pistil, a
seed vessel with ovules, and, growing to it, another bud of which the
structure was indistinct. 2nd flower, large; calyx of seven
divisions, one being transformed into a bud with bract, but much
smaller than the other; corolla large but cleft along the top; six
stamens with anthers, pistil, and seed-vessel. 3rd flower, large;
six-cleft calyx, cleft corolla, with six stamens, pistil, and
seed-vessel, with a second pistil half unfolded at its apex. 4th
flower, large; divided along the top, six stamens. 5th flower, large;
corolla divided into three parts, six stamens. 6th flower, large;
corolla cleft, calyx six cleft, the rest of the flower normal. 7th,
and all succeeding flowers, normal.
While this chapter is under revision, another noteworthy illustration
has been furnished to me by a wall-trained pear tree which was
covered in the spring by luxuriant "foreright" shoots. As I learned
from the gardener, it was pruned just as the fruit was setting. A
large excess of sap was thus thrown into other branches, with the
result that in a number of them the young pears were made monstrous
by reversion. In some cases, instead of the dried up sepals at the
top of the pear, there were produced good sized leaves; and in other
cases the seed-bearing core of the pear was transformed into a growth
which protruded through the top of the pear in the shape of a new
shoot.
[32] In partial verification, Mr. Tansley writes:--"Prof. Klebs of Basel
has shown that in _Hydrodictyon_, gametes can only be produced by the
cells of a net when these are above a certain size and age; and then
only under conditions unfavourable to growth, such as a feeble light
or poverty of nutritive inorganic salts or absence of oxygen, or a
low temperature in the water containing the plant. The presence of
organic substances, especially sugar, also acts as a stimulus to the
formation of gametes, and this is also the case in _Vaucheria_. Many
other _Algæ_ produce gametes mainly at the end of the vegetative
season, when food is certainly difficult to obtain in their natural
habitat, and we may well suppose that their assimilative power is
waning. Where, however, as is the case in _Vaucheria_, the plant
depends for propagation mainly on the production of fertilized eggs,
we find the sexual organs often produced in conditions very
favourable to vegetative growth, in opposition to those cases such as
_Hydrodictyon_, where the chief means of propagation is by zoospores.
So that side by side with, and to some extent obscuring, the
principle developed above we have a clear adaptation of the
production of reproductive cells to the special circumstances of the
case."
[33] This establishment by survival of the fittest of reproductive
processes adapted to variable conditions, is indirectly elucidated by
the habits of salmon. As salmon thrive in the sea and fall out of
condition in fresh water (having during their sea-life not exercised
the art of catching fresh-water prey), the implication is that the
species would profit if all individuals ran up the rivers just before
spawning time in November. Why then do most of them run up during
many preceding months? Contemplation of the difficulties which lie in
the way to the spawning grounds, will, I think, suggest an
explanation. There are falls to be leaped and shallow rapids to be
ascended. These obstacles cannot be surmounted when the river is low.
A fish which starts early in the season has more chances of getting
up the falls and the rapids than one which starts later; and, out of
condition as it will be, may spawn, though not well. On the other
hand, one which starts in October, if floods occur appropriately, may
reach the upper waters and then spawn to great advantage; but in the
absence of adequate rains it may fail altogether to reach the
spawning grounds. Hence the species profits by an irregularity of
habits adapted to meet irregular contingencies.
[34] I owe to Mr. (now Sir John) Lubbock an important confirmation of this
view. After stating his belief that between Crustaceans and Insects
there exists a physiological relation analogous to that which exists
between water vertebrata and land-vertebrata, he pointed out to me
that while among Insects there is a definite limit of growth, and an
accompanying definite commencement of reproduction, among
Crustaceans, where growth has no definite limit, there is no definite
relation between the commencement of reproduction and the decrease or
arrest of growth.
[35] While this chapter is passing through the press, I learn from Mr.
White Cooper, that not only are near sight, long sight, dull sight,
and squinting, hereditary; but that a peculiarity of vision confined
to one eye is frequently transmitted: re-appearing in the same eye in
offspring.
[36] An instance here occurs of the way in which those who are averse to a
conclusion will assign the most flimsy reasons for rejecting it.
Rather than admit that the eyes of these creatures living in darkness
have disappeared from lack of use, some contend that such creatures
would be liable to have their eyes injured by collisions with
objects, and that therefore natural selection would favour those
individuals in which the eyes had somewhat diminished and were least
liable to injury: the implication being that the immunity from the
inflammations due to injuries would be so important a factor in life
as to cause survival. And this is argued in presence of the fact that
one of the most conspicuous among these blind cave-animals is a
cray-fish, and that the cray-fish in its natural habitat is in the
habit of burrowing in the banks of rivers holes a foot or more deep,
and has its eyes exposed to all those possible blows and frictions
which the burrowing involves!
[37] In addition to the numerous illustrations given by Mr. Sedgwick, here
is one which Colonel A. T. Fraser published in _Nature_ for Nov. 9,
1893, concerning two Hindoo dwarfs:--"In speech and intelligence the
dwarfs were indistinguishable from ordinary natives of India. From an
interrogation of one of them, it appeared that he belonged to a
family all the male members of which have been dwarfs for several
generations. They marry ordinary native girls, and the female
children grow up like those of other people. The males, however,
though they develop at the normal rate until they reach the age of
six, then cease to grow, and become dwarfs."
[38] This remarkable case appears to militate against the conclusion,
drawn a few pages back, that the increase of a peculiarity by
coincidence of "spontaneous variations" in successive generations, is
very improbable; and that the special superiorities of musical
composers cannot have thus arisen. The reply is that the extreme
frequency of the occurrence among so narrow a class as that of
musical composers, forbids the interpretation thus suggested.
[39] I omitted to name here a cause which may be still more potent in
producing irregularity in the results of cousin-marriages. So far as
I can learn, no attempt has been made to distinguish between such
results as arise when the related parents from whom the cousins
descend are of the same sex and those which arise when they are of
different sexes. In the one case two sisters have children who
intermarry; and in the other case a brother and a sister have
children who intermarry. The marriages of cousins in these two cases
may be quite dissimilar in their results. If there is a tendency to
limitation of heredity by sex--if daughters usually inherit more from
the mother than sons do, while sons inherit more from the father than
from the mother, then two sisters will on the average of cases be
more alike in constitution than a sister and a brother. Consequently
the descendants of two sisters will differ less in their
constitutions than the descendants of a brother and a sister; and
marriage in the first case will be more likely to prove injurious
from absence of dissimilarity in the physiological units than
marriage in the second. My own small circle of friends furnishes
evidence tending to verify this conclusion. In one instance two
cousins who intermarried are children of two sisters, and they have
no offspring. In another the cousins who intermarried are children of
two brothers, and they have no offspring. In the third case the
cousins were descendants of two brothers and only one child resulted.
[40] _A propos_ of this sentence one of my critics writes:--"I cannot find
in this book the statement as first made that the 'life of an
individual is maintained by the unequal and ever-varying actions of
incident forces on its different parts.' Recent physiological work
offers a startling example of the statement."
To the question contained in the first sentence the answer is that I
have not made the statement in the above words, but that it is
implied in the chapter entitled "The Degree of Life varies as the
Degree of Correspondence," and more especially in § 36, which,
towards its close, definitely involves the statement. The verifying
evidence my critic gives me is this:--
"Prof. Sherrington has shown that if the sensory roots of the spinal
nerves are cut one by one there is at first no general effect
produced. That is to say, the remainder of the nervous system
continues to function as before. This condition (lack of general
effect) persists until about six pairs have been cut. With the
severance of the seventh pair, however, the whole central nervous
system ceases to function, so that stimulation of intact sensory
nerves produces no reflex action. After a variable period, but one of
many hours duration, the power of functioning is recovered. That is
to say, if the sensory impulses (from the skin, &c.) reaching the
central nervous system are rapidly reduced in amount, there comes a
point where those remaining do not suffice to keep the structure
'awake.' After a time, however, it adjusts itself to work with the
diminished supply. Similarly Strumpell describes the case of a boy
'whose sensory inlets were all paralyzed except one eye and one ear.'
When these were closed he instantly fell asleep."
[41] Fifty years before the discovery of the Röntgen rays and those
habitually emanating from uranium, it had been observed by Moser that
under certain conditions the surfaces of metals receive permanent
impressions from appropriate objects placed upon them. Such facts
show that the molecules of substances propagate in all directions
special ethereal undulations determined by their special
constitutions.
[42] This classification, and the three which follow it, I quote
(abridging some of them) from Prof. Agassiz's "Essay on
Classification."
[43] For explanations, see "Illogical Geology," _Essays_, Vol. I. How much
we may be misled by assuming that because the remains of creatures of
high types have not been found in early strata, such creatures did
not exist when those strata were formed, has recently (1897) been
shown by the discovery of a fossil Sea-cow in the lower Miocene of
Hesse-Darmstadt. The skeleton of this creature proves that it
differed from such Sirenian mammals as the existing Manatee only in
very small particulars: further dwindling of disused parts being an
evident cause. The same is true as regards, now, we consider that
since the beginning of Miocene days this aberrant type of mammal has
not much increased its divergence from the ordinary mammalian type;
if we then consider how long it must have taken for this large
aquatic mammal (some eight or ten feet long) to be derived by
modification from a land-mammal; and if then we contemplate the
probable length of the period required for the evolution of that
land-mammal out of a pre-mammalian type; we seem carried back in
thought to a time preceding any of our geologic records. We are shown
that the process of organic evolution has most likely been far slower
than is commonly supposed.
[44] Since this passage was written, in 1863, there has come to light much
more striking evidence of change from a more generalized to a less
generalized type during geologic time. In a lecture delivered by him
in 1876, Prof. Huxley gave an account of the successive modifications
of skeletal structure in animals allied to the horse. Beginning with
the _Orohippus_ of the Eocene formation, which had four complete toes
on the front limb and three toes on the hind limb, he pointed out the
successive steps by which in the _Mesohippus_, _Miohippus_,
_Protohippus_, and _Pliohippus_, there was a gradual approach to the
existing horse.
[45] Several of the arguments used in this chapter and in that which
follows it, formed parts of an essay on "The Development Hypothesis,"
originally published in 1852.
[46] _Studies from the Morphological Laboratory in the University of
Cambridge_, vol. vi, p. 84.
[47] _Ibid._, p. 81.
[48] _Studies from the Morphological Laboratory in the University of
Cambridge_, vol. vi, p. 89.
[49] Early in our friendship (about 1855) Prof. Huxley expressed to me his
conviction that all the higher articulate animals have twenty
segments or somites. That he adhered to this view in 1880, when his
work on _The Crayfish_ was published, is shown by his analysis there
given of the twenty segments existing in this fluviatile crustacean;
and adhesion to it had been previously shown in 1877, when his work
on _The Anatomy of Invertebrated Animals_ was published. On p. 398 of
that work he writes:--"In the abdomen there are, at most, eleven
somites, none of which, in the adult, bear ambulatory limbs. Thus,
assuming the existence of six somites in the head, the normal number
of somites in the body of insects will be twenty, as in the higher
_Crustacea_ and _Arachnida_." To this passage, however, he puts the
note:--"It is open to question whether the podical plates represent a
somite; and therefore it must be recollected that the total number of
somites, the existence of which can be actually demonstrated in
insects, is only seventeen, viz., four for the head, three for the
thorax, and ten for the abdomen." I have changed the number twenty,
which in the original edition occurred in the text, to the number
seventeen in deference to suggestions made to me; though I find in
Dr. Sharp's careful and elaborate work on the _Insecta_, that
Viallanes and Cholodkovsky agree with Huxley in believing that there
are six somites in the insect-head. The existence of a doubt on this
point, however, does not essentially affect the argument, since there
is agreement among morphologists respecting the _constancy_ of the
total number of somites in insects.
[50] To avoid circumlocution I let these words stand, though they are not
truly descriptive; for the prosperity of imported species is largely,
if not mainly, caused by the absence of those natural enemies which
kept them down at home.
[51] While these pages are passing through the press (in 1864), Dr. Hooker
has obliged me by pointing out that "plants afford many excellent
examples" of analogous transitions. He says that among true "water
plants," there are found, in the same species, varieties which have
some leaves submerged and some floating; other varieties in which
they are all floating; and other varieties in which they are all
submerged. Further, that many plants characterized by floating
leaves, and which have all their leaves floating when they grow in
deeper water, are found with partly aerial leaves when they grow in
shallower water; and that elsewhere they occur in almost dry soil
with all their leaves aerial.
[52] It will be seen that the argument naturally leads up to this
expression--Survival of the Fittest--which was here used for the
first time. Two years later (July, 1866) Mr. A. R. Wallace wrote to
Mr. Darwin contending that it should be substituted for the
expression "Natural Selection." Mr. Darwin demurred to this proposal.
Among reasons for retaining his own expression he said that I had
myself, in many cases, preferred it--"continually using the words
Natural Selection." (_Life and Letters_, &c., vol. III, pp. 45-6.)
Mr. Darwin was quite right in his statement, but not right in the
motive he ascribed to me. My reason for frequently using the phrase
"Natural Selection," after the date at which the phrase "Survival of
the Fittest" was first used above, was that disuse of Mr. Darwin's
phrase would have seemed like an endeavour to keep out of sight my
own indebtedness to him, and the indebtedness of the world at large.
The implied feeling has led me ever since to use the expressions
Natural Selection and Survival of the Fittest with something like
equal frequency.
[53] I am indebted to Mr. [now Sir W.] Flower for the opportunity of
examining the many skulls in the Museum of the College of Surgeons
for verification of this. Unfortunately the absence, in most cases,
of some or many teeth, prevented me from arriving at that specific
result which would have been given by weighing a number of the under
jaws in each race. Simple inspection, however, disclosed a
sufficiently-conspicuous difference. The under jaws of Australians
and Negroes, when collated with those of Englishmen, were visibly
larger, not only relatively but absolutely. One Australian jaw only
seemed about of the same size as an average English jaw; and this
(probably the jaw of a woman), belonging as it did to a smaller
skull, bore a greater ratio to the whole body of which it formed
part, than did an English jaw of the same actual size. In all the
other cases, the under jaws of these inferior races (containing
larger teeth than our own) were _absolutely_ more massive than our
own--often exceeding them in all dimensions; and _relatively_ to
their smaller skeletons were much more massive. Let me add that the
Australian and Negro jaws are thus strongly contrasted, not with all
British jaws, but only with the jaws of the civilized British. An
ancient British skull in the collection possesses a jaw almost or
quite as massive as those of the Australian skulls. All this is in
harmony with the alleged relation between greater size of jaws and
greater action of jaws, involved by the habits of savages.
[In 1891 Mr. F. Howard Collins carefully investigated this matter:
measuring ten Australian, ten Ancient British, and ten recent English
skulls in the College of Surgeons Museum. The result proved an
absolute difference of the kind above indicated, and a far greater
relative difference. To ascertain this last a common standard of
comparison was established--an equal size of skull in all the cases;
and then when the relative masses or cubic sizes of the jaws were
calculated, the result which came out was this:--Australian jaw,
1948; Ancient British jaw, 1135; Recent English jaw, 1030. "Hence,"
in the words of Mr. Collins, "the mass of the Recent English jaw is,
roughly speaking, half that of the Australian relatively to that of
the skull, and a ninth less than that of the Ancient British." He
adds verifying evidence from witnesses who have no hypothesis to
support--members of the Odontological Society. The Vice-President,
Mr. Mummery, remarks of the Australians that "the jaw-bones are
powerfully developed, and large in proportion to the cranium."]
[54] As bearing on the question of the varieties of Man, let me here refer
to a paper on "The Origin of the Human Races" read before the
Anthropological Society, March 1st, 1864, by Mr. Alfred Wallace. In
this paper, Mr. Wallace shows that along with the attainment of that
intelligence implied by the use of implements, clothing, &c., there
arises a tendency for modifications of brain to take the place of
modifications of body: still, however, regarding the natural
selection of spontaneous variations as the cause of the
modifications. But if the foregoing arguments be valid, natural
selection here plays but the secondary part of furthering the
adaptations otherwise caused. It is true that, as Mr. Wallace argues,
and as I have myself briefly indicated (see _Westminster Review_, for
April, 1852, pp. 496-501), the natural selection of races leads to
the survival of the more cerebrally-developed, while the less
cerebrally-developed disappear. But though natural selection acts
freely in the struggle of one society with another; yet, among the
units of each society, its action is so interfered with that there
remains no adequate cause for the acquirement of mental superiority
by one race over another, except the inheritance of
functionally-produced modifications.
[55] _Darwin and after Darwin_, Part II, p. 99.
[56] _Essays upon Heredity_, vol. i, p. 90.
[57] In a letter published by Dr. Romanes in _Nature_, for April 26, 1894,
he alleges three reasons why "as soon as selection is withdrawn from
an organ the _minus_ variations of that organ outnumber the _plus_
variations." The first is that "the survival-mean must descend to the
birth-mean." The interpretation of this is that if the members of a
species are on the average born with an organ of the required size,
and if they are exposed to natural selection, then those in which the
organ is relatively small will some of them die, and consequently the
mean size of the organ at adult age will be greater than at birth.
Contrariwise, if the organ becomes useless and natural selection does
not operate on it, this difference between the birth-mean and the
survival-mean disappears. Now here, again, the _plus_ variations and
their effects are ignored. Supposing the organ to be useful, it is
tacitly assumed that while _minus_ variations are injurious, _plus_
variations are not injurious. This is untrue. Superfluous size of an
organ implies several evils:--Its original cost is greater than
requisite, and other organs suffer; the continuous cost of its
nutrition is unduly great, involving further injury; it adds
needlessly to the weight carried and so again is detrimental; and
there is in some cases yet a further mischief--it is in the way.
Clearly, then, those in which _plus_ variations of the organ have
occurred are likely to be killed off as well as those in which
_minus_ variations have occurred; and hence there is no proof that
the survival-mean will exceed the birth-mean. Moreover the assumption
has a fatal implication. To say that the survival-mean of an organ is
greater than the birth-mean is to say that the organ is greater _in
proportion to other organs_ than it was at birth. What happens if
instead of one organ we consider all the organs? If the survival-mean
of a particular organ is greater than its birth-mean, the survival
mean of each other organ must also be greater. Thus the proposition
is that every organ has become larger in relation to every other
organ!--a marvellous proposition. I need only add that Dr. Romanes'
inferences with respect to the two other causes--atavism and failing
heredity--are similarly vitiated by ignoring the plus variations and
their effects.
[58] _Westminster Review_, January, 1860. See also _Essays, &c._, vol. i,
p. 290.
[59] "On Orthogenesis and the Impotence of Natural Selection in
Species-Formation," pp. 2, 19, 22, 24.
[60] Address to Plymouth Institution, at opening of Session 1895-6.
[61] _Westminster Review_, April, 1857. "Progress: its Law and Cause." See
also _Essays_, vol. i.
[62] It may be needful to remark, that by the proposed expression it is
intended to define--not Life in its essence; but, Life as manifested
to us--not Life as a _noumenon_: but, Life as a _phenomenon_. The
ultimate mystery is as great as ever: seeing that there remains
unsolved the question--What _determines_ the co-ordination of
actions?
[63] _Prin. of Phys._, 2nd edit., p. 77.
[64] _Ibid._, 3rd edit., p 249.
[65] _Ibid._, p. 124.
[66] Agassiz and Gould, p. 274.
[67] _Prin. of Phys._, 3rd edit., p. 964.
[68] "Parthenogenesis," p. 8.
[69] _Prin. of Phys._, p. 92.
[70] _Ibid._, p. 93.
[71] _Ibid._, p. 917.
[72] "A General Outline of the Animal Kingdom." By Prof. T. R. Jones, F.
G. S., p. 61.
[73] Carpenter.
[74] _Prin. of Phys._, p. 873.
[75] _Ibid._, p. 203.
[76] _Ibid._, p. 209.
[77] _Ibid._, p. 249.
[78] _Ibid._, p. 249.
[79] _Ibid._, p. 250.
[80] _Prin. of Phys._, p. 256.
[81] _Ibid._, p. 212.
[82] _Ibid._, p. 266.
[83] _Prin. of. Phys._, p. 267.
[84] _Ibid._, p. 276.
[85] _Ibid._, 2nd edit., p. 115.
[86] _Prin. of Phys._, p. 954.
[87] _Ibid._, p. 958.
[88] _Ibid._, p. 688.
[89] _Ibid._, p. 958.
[90] "A General Outline of the Animal Kingdom." By Professor T. R. Jones,
p. 61.
[91] _Prin. of Phys._, p. 907.
[92] Should it be objected that in the higher plants the sperm-cell and
germ-cell differ, though no distinct co-ordinating system exists, it
is replied that there _is_ co-ordination of actions, though of a
feeble kind, and that there must be some agency by which this is
carried on.
[93] It is a significant fact that amongst the dioecious invertebrata,
where the nutritive system greatly exceeds the other systems in
development, the female is commonly the largest, and often greatly
so. In some of the Rotifera the male has no nutritive system at all.
See _Prin. of Phys._, p. 954.
[94] _Prin. of Phys._, p. 908.
[95] "Parthenogenesis," pp. 66, 67.
[96] "Lectures on Animal Chemistry." By Dr. Bence Jones. _Medical
Times_, Sept. 13th, 1851. See also _Prin. of Phys._, p. 171.
[97] _Cyclopædia of Anatomy and Physiology_, Vol. IV, p. 506.
[98] From a remark of Drs. Wagner and Leuckart this chemical evidence
seems to have already suggested the idea that the sperm-cell becomes
"metamorphosed into the central parts of the nervous system." But
though they reject this assumption, and though the experiments of Mr.
Newport clearly render it untenable, yet none of the facts latterly
brought to light conflict with the hypothesis that the sperm-cell
contains unorganized co-ordinating matter.
[99] Quain's _Elements of Anatomy_, p. 672.
[100] The maximum weight of the horse's brain is 1 lb. 7 ozs.; the human
brain weighs 3 lbs., and occasionally as much as 4 lbs.; the brain of
a whale, 75 feet long, weighed 5 lbs. 5 ozs.; and the elephant's
brain reaches from 8 lbs. to 10 lbs. Of the whale's fertility we know
nothing; but the elephant's quite agrees with the hypothesis. The
elephant does not attain its full size until it is thirty years old,
from which we may infer that it arrives at a reproductive age later
than man does; its period of gestation is two years, and it produces
one at a birth. Evidently, therefore, it is much less prolific than
man. See Müller's _Physiology_ (Baly's translation), p. 815, and
Quain's _Elements of Anatomy_, p. 671.
[101] That the size of the nervous system is the measure of the ability to
maintain life, is a proposition that must, however, be taken with
some qualifications. The ratio between the amounts of gray and white
matter present in each case is probably a circumstance of moment.
Moreover, the temperature of the blood may have a modifying
influence; seeing that small nervous centres exposed to rapid
oxidation will be equivalent to larger ones more slowly oxidized.
Indeed, we see amongst mankind, that though, in the main, size of
brain determines mental power, yet temperament exercises some
control. There is reason to think, too, that certain kinds of nervous
action involve greater consumption of nervous tissue than others; and
this will somewhat complicate the comparisons. Nevertheless, these
admissions do not affect the generalization as a whole, but merely
prepare us to meet with minor irregularities.
[102] Let me here note in passing a highly significant implication. The
development of nervous structures which in such cases take place,
cannot be limited to the finger-ends. If we figure to ourselves the
separate sensitive areas which severally yield independent feelings,
as constituting a network (not, indeed, a network sharply marked out,
but probably one such that the ultimate fibrils in each area intrude
more or less into adjacent areas, so that the separations are
indefinite), it is manifest that when, with exercise, the structure
has become further elaborated, and the meshes of the network smaller,
there must be a multiplication of fibres communicating with the
central nervous system. If two adjacent areas were supplied by
branches of one fibre, the touching of either would yield to
consciousness the same sensation: there could be no discrimination
between points touching the two. That there may be discrimination,
there must be a distinct connection between each area and the tract
of grey matter which receives the impressions. Nay more, there must
be, in this central recipient-tract, an added number of the separate
elements which, by their excitements, yield separate feelings. So
that this increased power of tactual discrimination implies a
peripheral development, a multiplication of fibres in the
trunk-nerve, and a complication of the nerve-centre. It can scarcely
be doubted that analogous changes occur under analogous conditions
throughout all parts of the nervous system--not in its sensory
appliances only, but in all its higher co-ordinating appliances, up
to the highest.
[103] _Essays upon Heredity_, p. 87.
[104] _Les Maladies des Vers à soie_, par L. Pasteur, Vol. I, p. 39.
[105] Curiously enough, Weismann refers to, and recognizes, syphilitic
infection of the reproductive cells. Dealing with Brown-Séquard's
cases of inherited epilepsy (concerning which, let me say, that I do
not commit myself to any derived conclusions), he says:--"In the case
of epilepsy, at any rate, it is easy to imagine [many of Weismann's
arguments are based on things 'it is easy to imagine'] that the
passage of some specific organism through the reproductive cells may
take place, as in the case of syphilis" (p. 82). Here is a sample of
his reasoning. It is well known that epilepsy is frequently caused by
some peripheral irritation (even by the lodging of a small foreign
body under the skin), and that, among peripheral irritations causing
it, imperfect healing is one. Yet though, in Brown-Séquard's cases, a
peripheral irritation caused in the parent by local injury was the
apparent origin, Weismann chooses gratuitously to assume that the
progeny were infected by "some specific organism," which produced the
epilepsy! And then though the epileptic virus, like the syphilitic
virus, makes itself at home in the egg, the parental protoplasm is
not admitted!
[106] _Philosophical Transactions of the Royal Society for the Year 1821_,
Part I, pp. 20-24.
[107] It will, I suppose, be said that the non-inheritance of mutilations
constitutes evidence of the kind here asked for. The first reply is
that the evidence is conflicting, as it may well be. It is forgotten
that to have valid evidence of non-inheritance of mutilations, it is
requisite that both parents shall have undergone mutilation, and that
this does not often happen. If they have not, then, assuming the
inheritableness of mutilations, there would, leaving out other
causes, be an equal tendency to appearance and non-appearance of the
mutilation in offspring. But there is another cause--the tendency to
reversion, which ever works in the direction of cancelling individual
characters by the return to ancestral characters. So that even were
the inheritance of mutilations to be expected (and for myself I may
say that its occurrence surprises me), it could not be reasonably
looked for as more than exceptional: there are two strong
countervailing tendencies. But now, in the second place, let it be
remarked that the inheritance or non-inheritance of mutilations is
beside the question. The question is whether modifications of parts
produced by modifications of functions are inheritable or not. And
then, by way of disproof of their inheritableness, we are referred to
cases in which the modifications of parts are not produced by
modifications of functions, but are otherwise produced!
[108] See _First Principles_, Part II, Chap. XXII, "Equilibration."
[109] _Principles of Biology_, § 46, (No. 8. April, 1863).
[110] _Ibid._ This must not be understood as implying that while the mass
increases as the cubes, the _quantity of motion_ which can be
generated increases only as the squares; for this would not be true.
The quantity of motion is obviously measured, not by the sectional
areas of the muscles alone, but by these multiplied into their
lengths, and therefore increases as the cubes. But this admission
leaves untouched the conclusion that the ability to _bear stress_
increases only as the squares; and thus limits the ability to
generate motion, by relative incoherence of materials.
[111] _The Transactions of the Linnæan Society of London_, Vol. XXII, p.
215. The estimate of Reaumur, cited by Kirby and Spence, is still
higher--"in five generations one Aphis may be the progenitor of
5,904,900,000 descendants; and that it is supposed that in one year
there may be twenty generations." (_Introduction to Entomology_, Vol.
I, p. 175)
[112] _A Manual of the Anatomy of Invertebrated Animals_, by T. H. Huxley,
p. 206.
[113] Respecting the _Eloidea_ I learn that in 1879--thirty years after it
had become a pest--one solitary male plant was found in a pond near
Edinburgh; but "in an exhaustive inquiry on the plant made by Dr.
Groenland, of Copenhagen, he could find no trace of any male
specimens having been found in Europe other than the Scotch." In
waters from which the _Eloidea_ has disappeared, it seems to have
done so in consequence of the growth of an _Alga_, which has produced
turbid water unfavourable to it. That is to say, the decreased
multiplication of somatic cells in some cases, is not due to any
exhaustion, but is caused by the rise of enemies or adverse
conditions; as happens generally with introduced species of plants
and animals which multiply at first enormously, and then, without any
loss of reproductive power, begin to decrease under the antagonizing
influences which grow up.
[114] _A Text Book of Human Physiology._ By Austin Flint, M.D., LL.D.
Fourth edition. New York: D. Appleton & Co. 1888. Page 797.
[115] This supposition I find verified by Mr. A. S. Packard in his
elaborate monograph on "The Cave Fauna of North America, &c.," as
also in his article published in the _American Naturalist_,
September, 1888; for he there mentions "variations in _Pseudotremia
cavernarum_ and _Tomocerus plumbeus_, found living near the entrance
to caves in partial daylight." The facts, as accumulated by Mr.
Packard, furnished a much more complete answer to Prof. Lankester
than is above given, as, for example, the "blindness of _Neotoma_, or
the Wood-Rat of Mammoth Cave." It seems that there are also "cave
beetles, with or without rudimentary eyes," and "eyeless spiders" and
Myriapods. And there are insects, as some "species of Anophthalmus
and Adelops, whose larvæ are lacking in all traces of eyes and optic
nerves and lobes." These instances cannot be explained as sequences
of an inrush of water carrying with it the remote ancestors, some of
which did not find their way out; nor can others of them be explained
by supposing an inrush of air, which did the like.
[116] See "Social Organism" in _Westminster Review_ for January, 1860; also
_Principles of Sociology_, § 247.
[117] _Contemporary Review_, September, 1893.
[118] _Evolution of Sex_, p. 50.
[119] _Souvenirs Entomologiques_, 3^{me} Série, p. 328.
[120] _Natural History of Bees_, new ed., p. 33.
[121] _Origin of Species_, 6th ed., p. 232.
[122] _Contemporary Review_, September, 1893, p. 333.
[123] _The Entomologist's Monthly Magazine_, March, 1892, p. 61.
[124] Perhaps it will be alleged that nerve-matter is costly, and that this
minute economy might be of importance. Anyone who thinks this will no
longer think it after contemplating a litter of half-a-dozen young
rabbits (in the wild rabbit the number varies from four to eight);
and on remembering that the nerve-matter contained in their brains
and spinal cords, as well as the materials for building up the bones,
muscles, and viscera of their bodies, has been supplied by the doe in
the space of a month; at the same time that she has sustained herself
and carried on her activities: all this being done on relatively poor
food. Nerve-matter cannot be so very costly then.
[125] _Loc. cit._, p. 318.
[126] _The Germ Plasm_, p. 54.
[127] While Professor Weismann has not dealt with my argument derived from
the distribution of discriminativeness on the skin, it has been
criticized by Mr. McKeen Cattell, in the last number of _Mind_
(October, 1893). His general argument, vitiated by extreme
misconceptions, I need not deal with. He says:--"Whether changes
acquired by the individual are hereditary, and if so to what extent,
is a question of great interest for ethics no less than for biology.
But Mr. Spencer's application of this doctrine to account for the
origin of species [!] simply begs the question. He assumes useful
variations [!]--whether of structure or habit is immaterial--without
attempting to explain their origin": two absolute misstatements in
two sentences! The only part of Mr. Cattell's criticism requiring
reply is that which concerns the "sensation-areas" on the skin. He
implies that since Weber, experimental psychologists have practically
set aside the theory of sensation areas: showing, among other things,
that relatively great accuracy of discrimination can be quickly
acquired by "increased interest and attention.... Practice for a few
minutes will double the accuracy of discrimination, and practice on
one side of the body is carried over to the other." To me it seems
manifest that "increased interest and attention" will not enable a
patient to discriminate two points where a few minutes before he
could perceive only one. That which he can really do in this short
time is to learn to discriminate between the _massiveness of a
sensation_ produced by two points and the massiveness of that
produced by one, and to _infer_ one point or two points accordingly.
Respecting the existence of sensation-areas marked off from one
another, I may, in the first place, remark that since the eye
originates as a dermal sac, and since its retina is a highly
developed part of the sensitive surface at large, and since the
discriminative power of the retina depends on the division of it into
numerous rods and cones, each of which gives a separate
sensation-area, it would be strange were the discriminative power of
the skin at large achieved by mechanism fundamentally different. In
the second place I may remark that if Mr. Cattell will refer to
Professor Gustav Retzius's _Biologische Untersuchungen_, New Series,
vol. iv (Stockholm, 1892), he will see elaborate diagrams of
superficial nerve-endings in various animals showing many degrees of
separateness. I guarded myself against being supposed to think that
the sensation-areas are sharply marked off from one another; and
suggested, contrariwise, that probably the branching
nerve-terminations intruded among the branches of adjacent
nerve-terminations. Here let me add that the intrusion may vary
greatly in extent; and that where the intruding fibres run far among
those of adjacent areas, the discriminativeness will be but small,
while it will be great in proportion as each set of branching fibres
is restricted more nearly to its own area. All the facts are
explicable on this supposition.
[128] To save space and exclude needless complication I have omitted these
passages from the preceding divisions of this appendix.
[129] Though Professor Weismann does not take up the challenge, Dr. Romanes
does. He says:--"When selection is withdrawn there will be no
excessive _plus_ variations, because so long as selection was present
the efficiency of the organ was maintained at its highest level: it
was only the _minus_ variations which were then eliminated"
(_Contemporary Review_, p. 611). In the first place, it seems to me
that the phrases used in this sentence beg the question. It says that
"the efficiency of the organ was maintained at its _highest_ level";
which implies that the highest level (tacitly identified with the
greatest size) is the best and that the tendency is to fall below it.
This is the very thing I ask proof of. Suppose I invert the idea and
say that the organ is maintained at its right size by natural
selection, because this prevents increase beyond the size which is
best for the organism. Every organ should be in due proportion, and
the welfare of the creature as a whole is interfered with by excess
as well as by defect. It may be directly interfered with--as for
instance by too big an eyelid; and it may be indirectly interfered
with, where the organ is large, by needless weight and cost of
nutrition. In the second place the question which here concerns us is
not what natural selection will do with variations. We are concerned
with the previous question--What variations will arise? An organ
varies in all ways; and, unless reason to the contrary is shown, the
assumption must be that variations in the direction of increase are
as frequent and as great as those in the direction of decrease. Take
the case of the tongue. Certainly there are tongues inconveniently
large, and probably tongues inconveniently small. What reason have we
for assuming that the inconveniently small tongues occur more
frequently than the inconveniently large ones? None that I can see.
Dr. Romanes has not shown that when natural selection ceases to act
on an organ the _minus_ variations in each new generation will exceed
the _plus_ variations. But if they are equal the alleged process of
panmixia has no place.
[130] _The Variation of Animals and Plants under Domestication_, vol. ii,
p. 292.
[131] _Journal of the Anthropological Institute_ for 1885, p. 253.
[132] In "The All-Sufficiency of Natural Selection" (_Contemporary Review_,
Sept., 1893, p. 311), Professor Weismann writes:--"I have ever
contended that the acceptance of a principle of explanation is
justified, if it can be shown that without it certain facts are
inexplicable." Unless, then, Prof. Weismann can show that the
distribution of discriminativeness is otherwise explicable, he is
bound to accept the explanation I have given, and admit the
inheritance of acquired characters.
[133] Prof. Weismann is unaware that the view here ascribed to Roux,
writing in 1881, is of far earlier date. In the _Westminster Review_
for January, 1860, in an essay on "The Social Organism," I
wrote:--"One more parallelism to be here noted, is that the different
parts of a social organism, like the different parts of an individual
organism, compete for nutriment; and severally obtain more or less of
it according as they are discharging more or less duty." (See also
_Essays_, i, 290.) And then, in 1876, in _The Principles of
Sociology_, vol. i, § 247, I amplified the statement thus:--"All
other organs, therefore, jointly and individually, compete for blood
with each organ ... local tissue-formation (which under normal
conditions measures the waste of tissue in discharging function) is
itself a cause of increased supply of materials ... the resulting
competition, not between units simply, but between organs, causes in
a society, as in a living body, high nutrition and growth of parts
called into greatest activity by the requirements of the rest."
Though I did not use the imposing phrase
"intra-individual-selection," the process described is the same.
[134] _Proceedings of the Biological Society of Washington_, vol. ix.
[135] Romanes Lecture, p. 29.
[136] _Ibid._, p. 35.
[137] This interpretation harmonizes with a fact which I learn from Prof.
Riley, that there are gradations in this development, and that in
some species the ordinary neuters swell their abdomens so greatly
with food that they can hardly get home.